Phytotaxa 93 (1): 1–24 (2013) ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ PHYTOTAXA Copyright © 2013 Magnolia Press Article ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.93.1.1

The relationship and different C4 Kranz anatomy of Bassia eriantha and Bassia eriophora, two often confused Irano-Turanian and Saharo-Sindian species

HOSSEIN AKHANI1,* & ROXANA KHOSHRAVESH1 1 Department of Sciences, School of Biology and Center of Excellence in Phylogeny of Living Organisms, College of Sciences, University of Tehran, P. O. Box 14155-6455, Tehran, Iran * Corresponding author: [email protected]

Abstract

The circumscription and generic status of Bassia eriantha (≡ Londesia eriantha) and B. eriophora have often been confused in the literature. The reason is their extreme superficial similarity and phenology. In a multidisciplinary approach, we investigated both in the field, by cultivation in the laboratory, and performed anatomical, ultra-structural and molecular studies to clarify their and relationships. Both species are not only geographically and morphologically distinct by reliable and constant characters, but surprisingly also have different anatomical C4 Kranz types and occur in different clades using ITS nrDNA sequence analysis. Using the recent broad circumscription of the genus Bassia they belong to Bassia but in different clades. In spite of their rather well distinct geography, the two species are sympatric in the south-eastern Iran and south Afghanistan. Bassia eriantha as an Irano-Turanian species occurs disjunctly as Irano-Turanian enclave in south Sinai, Jordan and W Saudi Arabia as a second sympatric range co- occurring with B. eriophora.

Key words: , bundle sheath, C4 photosynthesis, Camphorosmeae, Chenopodiaceae, halophyte, phylogeny

Introduction

The increasing interest in the evolution of C4 photosynthesis pathway requires taxonomic studies within C4 taxa and their C3 relatives. Chenopodiaceae represents one of the major lineage of C4 diversity among

Eudicots having a multiple origin of C4 photosynthesis and a high morpho-anatomical variability (Akhani et al. 1997, Kadereit et al. 2003, Sage et al. 2011). In Eurasia this family is diversified extensively in the Middle East, Central (uplands) and especially Middle Asia (lowland around the Aral sea and Turan), and to some extent also in North and South African deserts, with a great success in formation of vegetation under harsh, dry and saline conditions and during very hot growing seasons. Camphorosmeae Endlicher (1837: 294), that is alternatively considered as subfamily Camphorosmioideae Scott (1978: 102), is a species-rich group which classification of this lineage has always been controversial in the literature of this family (Scott 1978, Kühn et al. 1993). A recent molecular investigation of this tribe (Kadereit & Freitag 2011) proposed a new classification in which Bassia Allioni (1766: 177) is widely circumscribed by including several mono- and oligo-specific genera such as Roth (1801: 307), Londesia Fischer & Meyer (1835: 40), Panderia Fischer & Meyer (1835: 46), Kirilowia Bunge (1843: 7) and Chenoleoides (Ulbrich 1934: 530) Botschantzev (1976: 1408). One of the species listed is Bassia eriophora (Schrader 1809: 86) Ascherson (in Schweinfurth 1867: 187) which was broadly circumscribed more recently (see below) and includes L. eriantha Fischer & Meyer (1835: 40) or B. eriantha (Fisch. & C. A. Mey.) Kuntze (1891: 546). Bassia eriophora was first described based on a cultivated plant under the genus Kochia (Schrader 1809). The origin of the seeds was doubtfully mentioned as Spain which, considering its known range, is unlikely. The illustration and description are fully informative and cannot be confused with any other plant. Kochia

Accepted by Duilio Iamonico: 12 Mar. 2013; published online in PDF: 11 Apr. 2013 1